Royal Ontario Museum
Domestication of the Carp
Cyprinus carpio L.
Eugene K. Balon
' / : "
Fig. 1 Coloration of Danube wild carp and reared offspring. Top: Danube wild carp,
female. 49 cm standard length, from lesser Danube above Kolarovo. Slovakia.
Centre: first generation of wild carp reared in ponds, cight-month-old fish, 15
cm long. Bottom: seven-year-old, "dwarfed" Danube wild carp, 15 cm long
(photo F. K. Balon).
ROYAL ONTARIO MUSEUM
eugene k.balon Domestication of the Carp
Cyprinus carpio L.
Royal Ontario Museum
Dr. A.D. Tushingham
Suggested citation: Roy.Ont.Mus. Life Sci. Misc. Pub.
Publication date: 29 April 1974
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e. K. balon is Associate Professor. Department of Zoology, University of
Guelph, and Research Associate. Department of Ichthyology and Herpetology,
Royal Ontario Museum.
©The Royal Ontario Museum, 1974
100 Queen's Park. Toronto, Canada
PRINTED AT THE BRYANT PRESS LIMITED
(Oryx, 1963. p.: 14)
"Preservation by domestication in an alien en-
viron/nent is far better than extinction in the
In spite of commonly held views that domesti-
cation preserves — it does not — it alters.
(R. L. Smith's
"'The Ecology of
Man: . . .", 1972, p.91)
"Domestication changed the life of the beast, the
character of the animal, and its anatomy and
The cover: Fishermen of the wild carp on the Danube River near Komarno one
hundred years ago (reprinted from an etching in O. Herman, A Magyar halaszat
konyve, Budapest, 1887).
Materials and Methods, 2
Paleogeographical Distribution of Carp, 4
Historical Evidence of Danubian Distribution, 5
Taxonomy and Evolution, 7
Dwarfed Wild Carp and their Viability, 8
Probable Origin of Domestication, 16
Roman Gourmets and First Carp in Captivity, 17
Fasting Monks Rediscover the Carp, 1 9
Rearing of Carp in Ponds and
First Reproduction in Captivity, 20
One More Origin of Domestication, 21
Consequences of Domestication, 22
Literature Cited, 27
Digitized by the Internet Archive
in 2012 with funding from
Royal Ontario Museum
Domestication of the Carp
Cyprinus carpio L.
Historical, zoogeographical, morphological, and physiological
information were utilized in explaining the origins and history
of domestication of the carp. The wild carp ancestor apparently
originated in central Asia and spread naturally east into China
and west as far as the Danube River. Evidence suggests that
the Romans first cultured carp collected from the Danube, and
the tradition of the piscinae was continued in monasteries
throughout the Middle Ages. A starvation experiment demon-
strated the hardiness of Danube wild carp and confirmed its
suitability for domestication. Domestication of the carp in
China was first independent of similar activities in Europe. The
strong evidence pointing to rheophilic Danube wild carp as
ancestral to modern domestic varieties suggests the importance
of preserving this natural population for further use.
Pour expliquer les origines et l'historique de la domestication
de la carpe on a mis en oeuvre differentes sources de rensei-
gnements; l'histoire. la zoogeographie, la morphologic et la
On trouve Tancetre de la carpe (a l'etat sauvage) apparem-
ment d'abord en Asie centrale, puis graduellement vers Test
jusqu'en Chine et vers l'ouest jusqu'au Danube. II y a tout
lieu de croire que e'etaient les Romains qui firent les premiers
l'elevagc de la carpe qu'ils prenaicnt du Danube, et on continua
la tradition des piscinae dans les monasteres pendant le Moyen
Age. On a tente de demontrer par des experiences avec la carpe
sauvage du Danube, la resistance de celle-ci a un regime de
jeune force, et ainsi on a meme prouve qu*on pourrait en faire
parfaitement un elevage systematique. L'elevage de la carpe en
Chine s'est developpe d*abord independemment de celui de
l'Europc. Les recherches fakes nous ont fourni la quasi-certi-
tude que la carpe sauvage reophile du Danube est bien 1'ancetre
des differentes cspeccs moderaes d'elevage, et ainsi clles mct-
tcnt meme au tout premier plan I'mterct a preserver cette popu-
lation naturelle pour des usages ulterieurs.
The wild carp (Cyprinn.s carpio carpio) is the predecessor of the domesti-
cated or pond carp about whose vicissitudes we have less information than
about the extinct auroch, the predecessor of cattle. An attempt is made, there-
fore, to document its evolutionary and domestic history.
During the past 400 years the domestic carp has been intensively cultured
in Europe and introduced into many countries around the world. The origin,
however, of this valuable commercial species has never been satisfactorily
explained. The writing of this paper was stimulated by recent publications
on the history of the carp in Europe (by Rudzihski, 1962, among others),
Australia (Butcher, 1967), and North America (among others, McCrimmon,
1968) that repeated old, unfounded ideas regarding the origin of this fish
and also omitted details of its prehistory and early history (Vooren, 1972).
Results of a number of studies (Balon and Misik, 1956; Misik, 1958;
Misik and Tuca, 1965; Balon, 1957, 1958a, 1958b; Bastl, 1961 ; Rudzihski,
1961 ) bearing on wild carp domestication are summarized here in an attempt
to relate biological results to facts gleaned from zoogeography, history, and
archaeology. As these data are meagre and their interpretation often ambigu-
ous or subjective, the origin of domestic carp may not be solved definitively
even in this essay. If my hypothesis, however, encourages more research and
assists archaeology and social anthropology to interpret their newly discov-
ered artifacts and data, the goal of this paper will have been accomplished.
An earlier version of this study appeared in the mimeographed "Works of
the Laboratory of Fishery Research in Bratislava*' (Balon, 1969). Presented
here is a revised and amended version, with a new section on an experiment
in starvation. The experiment was a result of an incidental activity, and
though lacking details of a strictly scientific study, is worthy of record.
Materials and Methods
The first numerous wild carp in the piedmont zone of the Danube River were
collected from a spawning school in May and June of 1955. Although I
sampled individual specimens a year earlier (Balon and Misik, 1956), the
intensified search resulted in finding the spawning school near the village of
Kolarovo. These fish formed the main material for the age and growth study
(Balon, 1957) and for the taxonomic revision (Misik, 1958). Although
initially added to the collection of the Laboratory of Fishery Research (Slo-
vak Agricultural Academy), most of them were later transferred to and made
a part of the collection of the Slovak National Museum in Bratislava. Some
specimens from successive spawning and developmental experiments (Balon.
1958a), pond culture (Bastl, 1961) and from the experiment in starvation
were also preserved and stored in both collections. These collections can now
be considered major retainers of preserved rheophilic wild carp of the Danube
The carp were kept in aquaria for an experiment in starvation (date of
hatching — 31 May 1956). Individual specimens were successively selected
for preservation or study of scales: first and second juveniles were preserved
on 13 October 1956 (standard length [SL] 44 and 61 mm, weight [w] 2 and
5 g respectively); third specimen, a mature female, was preserved 19 March
1959 (SL 128 mm. w 56 g); from the fourth specimen scales were studied
on 20 June 1958 (SL 129 mm, w 53 g) and the fish was preserved as a
mature male on 19 March 1959 (SL 144 mm, w 67 g); from the fifth speci-
men scales were studied on 20 June 1958 (when SL 120 mm, w 48 g) and
the fish was preserved as a mature male on 9 December 1960 (when SL 134
mm — the fish was unsuitable for most measurements because it jumped out
of the aquarium and was found dehydrated); from the sixth and seventh
specimen scales were studied on 13 October 1961 (SL 150 and 148 mm,
w 69 and 56 g respectively) and on 20 June 1958 (when SL 138 and 146
mm. w 63 and 80 g respectively), the sixth fish was preserved as a mature
female on 26 April 1962 (SL 148 mm, w 47 g), the seventh fish as a mature
female on 6 January 1963 (SL 148 mm, w 68 g).
The relative growth indices used in Table 1 were explained in an earlier
study (Balon, 1964, 1972). The identification of sex and usage of meristic
and morphometric characters (Table 2) is defined in detail in Misik's (1958)
study; the indices in Table 3 in Rudzinski's (1961) and Steffens' (1964)
Figures on the inside covers give some illustration of the extensive initial
studies of the wild carp. They are reprinted here primarily for the benefit of
the English language reader, and they explain the counts, measurements and
usage of terms.
INSIDE THE I RONT AND BACK COVERS
Front cover: a-g some developmental stages of the wild carp (from Balon,
1958a); a — embryo at time of hatching and its main respiratory organs —
ducti Cuvieri in anterior part of the yolk sac; c — feeding larva 12 days old
and 11 mm long with well developed (o) external lateral line sensors
(cupulae); E — full grown larva 20 days old and 18 mm long; F — the appear-
ance of first scales on a 15 mm long larva, and G — a 22 mm long juvenile.
h-m some characters of an adult wild carp (from Misik, 1958) ; H — the type
wild carp from the Danube River near Medvedovo (21.5.1954, Slovak
National Museum #171 ), I — its first gill arch and J — pharyngeal bone; K —
sketch indicating meristic characters; L, M — sketch indicating morphometric
characters (numbers refer to characters listed in Table 2).
Back cover: a — scaled morph, b — line or heavily scaled mirror morph,
c — lesser scaled mirror morph. and d — scaleless or leather morph of the
domestic carp (after Wunder, 1949); E — the type wild carp (from Balon,
1967b); F — the rate of growth of wild carp (from Balon. 1957) ;g — the wild
and H — the domestic carp (after Holcik and Hensel, 1972).
Paleugeographical Distribution of Carp
Studies proving the occurrence of the carp in Western Europe as early as the
Tertiary are based primarily on scales found in preglacial freshwater strata
in northern Germany (Nehring, 1883. after Zaunick. 1925) and at pile
dwelling sites in Switzerland. Ruetimeyer's (1860) claim that he found
scales of carp in remnants of pile dwellings was later contradicted by Forel
(1904). Forel based his opinion on the information of Studer (in Zaunick.
1925) who identified the scales as those of Abramis brama. Thus, not only
is the identity of preglacial remnants doubtful, but their age is unconfirmed.
There is no further true paleontological evidence. Consequently, the occur-
rence of wild carp in preglacial Europe is conjectural.
The most correct opinions seem to be those of authors (see in Misik,
1958) who considered Asia Minor and the area of the Caspian Sea to be the
origin of the wild carp. The primitive morph or subspecies of the wild carp,
Cyprinus carpio anatolicus Hanko, 1924, with the greatest number of
pharyngeal teeth, still occurs there. All morphs of the wild carp even today
are highly adaptable and almost always react to changing environmental
conditions by an abrupt modification of genotype (Dolzhenko, 1953; Bur-
makin, 1956; and others). Therefore, all morphs of wild carp may be con-
sidered to be evolutionarily young. That the primitive morph had started to
occupy streams peripheral to western Asia where it originated as early as the
late Pliocene is proved by remnants in "pontic" lacustrine strata (Borzenko,
1926; Hanko, 1932; Banarescu, 1960). However, it probably did not survive
the Pleistocene in that new area. Certain negative results support my hypothe-
sis. Had the wild carp lived in the Danube refuge in the Pleistocene it would
certainly have moved north to Scandinavia and the British Isles as conti-
nental glaciers retreated before the end of the period of interfluvial connec-
tions as did, for example, Abramis brama, Esox lucius, Rmilus rutilus, Scar-
dinius erythrophthalmus, Tinea tinea, Blicca bjoerkna, and Cobitis taenia
(Thiencmann, 1950). Carp also did not penetrate (e.g. via the Bering land
bridge) to North America in the Pleistocene, at a time when this connection
enabled Esox lucius to do so (Lindberg, 1962, Crossman and Harington,
1970). Therefore the carp likely had not reached the Far East by the end of
the Pliocene. Thus, the ancestors of recent carp probably evolved in the
Caspian area and spread from there to western Europe and to China as late
as the last postglacial period.
Except for the Danube River, the natural occurrence of carp in waters of
Europe as early as the beginning of the Christian era is not probable. Hence
Ausonius (a.d. 310-393; 1933) did not mention carp in the fauna of the
Rhine and Mosel rivers in the fourth century. Later records of sporadic
occurrence of carp in rivers may be explained by individuals having escaped
from ponds, though Dhigosz ( 1863-1887) did not mention the presence of
carp in Polish waters even as late as the 1 5th century.
The ancestor of contemporary wild carp evidently evolved in the area of
the Caspian Sea at the end of the Pleistocene. Under conditions of the post-
glacial thermal optimum some strains spread as far as the Black Sea area, the
Aral system, into eastern Asia and appeared in the Danube River system
about 8,000-10,000 years ago, that is, somewhat sooner than did the gold-
fish. Carassius auratus gibelio, which today penetrates the Danube River
(Hensel, 1971 ). The westward expansion of the range of that species was
probably similar to that of the wild carp (Balon, 1962. 1963a).
Jordan and Evermann (1896-1900, 1902) and Burns (1966) were thus
mistaken to consider the carp to be an Asian species that was introduced in
Europe before 1758 and was named after the Island of Cyprus, which they
considered to be the centre of its distribution. Also in error was Butcher
(1967) when stating. "'Whatever changes in the environment were brought
about following the introduction of this fish into Europe have been lost
apparently in antiquity."
According to ancient stories and myths, the Great Schutt Island (in the
upper part of the Middle Danube) was surrounded by a "great number of
golden carp (Cyprinus auratus) that enabled even the poorest people to make
a living; yes, there were times when the fishermen gave them away as gifts"
(translated from Khin. 1930). Then why did Jordan (and Evermann, 1896-
1902). Thienemann (1950). Maar (1960), Vooren (1972), and so many
others believe that the carp reached Rome via Greece from China? Possibly
even the carpio of Plinius (a.d. 24-79; 1635) was not an unknown seafish
but the wild carp; then, of course, the specific name could be from the Greek
root Karpos, meaning fruit.
HISTORICAL EVIDENCE OF DANUBIAN DISTRIBUTION
Natural occurrence of the wild carp in the Danube River is supported by the
evidence of Lconhardt ( 1906), who however hypothesized that the wild carp
arrived there from its northern distribution area before the formation of the
continental pack-ice. Leonhardt (1906) assented that names used for the
carp in local areas and the ultimate derivation of the scientific name can
tell much about the historical distribution of that fish; he advanced a probable
explanation of the generic and specific name of the carp. The generic name
is from latinized Greek and the species name comes from the Celtic col-
loquial name for the fish. "Kyprinos" or "Kyprianos" was the name given by
Aristotle (384-322 B.C.; 1862) to this fish and was probably derived from
"C\pris." a secondary name of the Goddess of Love, Aphrodite — perhaps
because the high fertility of the carp was known even then. Later the name
was latinized to "Cyprinus." probably by Pliny. Belon ( 1555) mentioned that
he encountered the carp in the land of the Etolic Greeks, who called it
The specific name, however, probably originated later and as "carpa"
appeared for the first time in the works of Cassiodorus (a.d. 490-585; 1626)
in the 6th century. The old Romans called it cyprinus, a name that spread
with the fish throughout Europe. The lack of indigenous local names for this
fish to the west and north of the Danube River may be considered to be
further evidence that the name originated there with the appearance of the
species. The Latin name of the fish is Celtic in origin and dates from the time
when Celtic tribes inhabited the present eastern Austrian and Czechoslovak-
Hungarian territory of the Danube River. Prom the ancient "charpho."
"carfo" and "charofo" the name gradually changed to the present "carpo."
Celts were certainly familiar with the carp at the time they settled in the
Danube area, from where the name spread with them or with later introduc-
tions of the fish. Everywhere the Celts lived, the root of the name "carp" oc-
curs — "carpe" in France; "carpio" or "carpione" in Italy; "carp" in England;
"carpe" in Spain; "Karpfen" in Germany; "karp" in Poland and Russia; and
"kapr" and "Kapor" in Czechoslovakia. In the region of the Danube River,
where it originated, however, this fish is called "ponty" (Hungarian),
"sharan" (Serbian), "saran" (Bulgarian), "crap," "saran," "ciortan,"
"ciuciu" or "olocari" (Rumanian), "husgun" (Turkish), "korop," "sharan,"
"podrojek" (Ukrainian), "sazan" (on the Volga River), and "kalynshyr"
Cassiodorus (a.d. 490-585; 1626) wrote about shipments of Danube
carp to Ravenna in Italy for the table of King Theodorus, and his note may
also serve as evidence of the Danubian origin of the western domestic carp.
Consequently, there is no basis for later opinions which were copied without
critical comment from one book to the next, that had the wild carp originate
in China or eastern Asia and suggested that European pond-culture was, so to
speak, the continuation of its earlier culture in China (Maar, 1960; Steffens,
1967). These concepts were supported by some authors (Dubravius, 1547;
von Hohberg, 1687) in the Middle Ages who wrote at a time when pond cul-
ture was spreading but completely misplaced the origin of the fish. They
found, correctly, that carp existed in European ponds before occurring in
rivers but erroneously included the Danube among the rivers. According to
von Hohberg ( 1687, p. 582) for example:
Die Sce-Karpfen (odcr die in den
Fliissen gcfangcn werden ) halt man
auch fur besser (und wiewol sie ge-
wohnlich in den Fliissen und Stro-
men nich wohnen) geschiehct es
doch vielmal (dass die abgebrochcn
Teichc ihre Fische mit samt dem
Wasser dahin uberlassen miissen)
wie man an dem Teyalluss spuhren
kan (dcr seine aus den Teichcn
enpfangenc Karpffen der March)
die March aber bei Toben der
Donau mitt-heilet (daher auch
zwischcn Prcssburg und Tbbcn vicl
Karpffen in der Donau zu finden
sind; sie gelangen zu grosscm Alter)
wiewol sic in vicr und fiinf Jahren
zu Speisc am dienlichsten.
The lake carp (or that caught in
rivers) is considered to be better,
and although it ordinarily does not
inhabit rivers and streams, it often
finds its way into rivers with water
from ponds with broken dams.
Carp, released from ponds, are
transferred first to the Dyje River
and then to the Morava River,
which joins the Danube at Dcvin.
In the Danube River between Brat-
islava and Devin occur many carp.
Individuals reach a great age, al-
though they are most edible when
they are 4 to 5 years old.
That individuals of the pond morph that escaped into rivers soon changed
from a short, scaleless morph into an elongate, scaled morph similar to the
wild carp supports von Hohberg's ( 1687) statement. A change in body shape
is also known (Tchen. 1956) in varieties of the goldfish (Carassius auratus
auratus). Although obser\ers in the Middle Ages considered the numerous
carp at the mouth of the Morava River near Devin to have escaped from
ponds, it was certainK the endemic, wild form of the carp that was abundant
in that part of the river, which has a large flood zone needed for successful
spawning of carp.
Taxonomy and Evolution
Recent native carp from the Danube River along the Czechoslovakia-
Hungary border are an ancient morph and ecologically isolated from escaped
or introduced pond carp ( Misik, 1 958 ) . But of all wild morphs of the carp —
from the Danube River. Aral Sea. and central and eastern Asia — that in the
Danube is nearest to the domestic carp (Balon, 1957, 1958a; Misik, 1958).
In other words, we consider the wild carp of the Danube River to be not a
form of pond carp which became feral after it was released into the river as is
generally accepted, but as a true wild morph related to the domestic pond carp
in a way similar to that of the ancestral auroch to cattle. In nature the wild
carp does not hybridize with the domestic carp. Domestic carp are never
found in schools of wild carp, and single, domestic carp caught within the
spawning grounds of wild carp were never ready to spawn. Domestic carp
which gradually change in rivers into a scaled, elongate morph similar to the
wild carp, were normally easily recognized by a notch and hump posterior
to the head; in native wild carp the transition between the top of the head
and the back (nape) is slightly if at all marked. Numerous breeding experi-
ments (Tuca. 1958; Bastl. 1961, 1962; Rudzinski, 1961, 1962; Misik and
Tuca. 1965) proved the genetic distinctness of this morph.
After a biometric study of wild Danube carp and after comparison with
wild carp from other regions. Misik (1958) demonstrated that wild carp
can be divided into three groups:
1. European wild carp (Cyprinus carpio carpio) represented by the popula-
tion of the Danube River system and designed as nominate subspecies;
2. Eastern Asian wild carp (Cyprinus carpio haeinatopterus) from Siberia
3. Wild carp "from the Aral Sea and from other central Asian regions which
in some ways are more closely related to the European ones, in others to east
Asian carp, which at the same time are mutually substantially difTerent ,,
(translated from Misik. 1958. p. 106).
Concerning the third group Misik (loc. cit.) concluded (my translation) :
"'Reasons for these convergent and divergent changes in morphological fea-
tures of the central Asian carp — in my opinion — are to be sought in the geo-
graphic conditions of the area, which may include specific peculiarities of the
geography of both Europe and eastern Asia. It is particularly those specific
characters that may show the orientation of the evolutionary divergence of
central Asian carp. That is why they may be considered as intcr-digitating.
allopatric populations, which — if they are geographically isolated from the
subspecies C. c. haematopterus and C. c. carpio — can be considered as sepa-
rate geographical morphs with numerous, ecological modifications." Misik's
divisions, however, may be interpreted also as follows:
The differentiation of the wild carp in the region of the Aral Sea and central
Asia was influenced by an earlier geological age; if central Asia is considered
to be the origin of the wild carp, descendants of that original strain had a
longer time, the entire Pleistocene, to evolve in isolation as compared to
carp that emigrated eastward and westward in the last postglacial period.
Even though strains later integrated, more morphs may exist today in west-
ern and central Asia than elsewhere. Regarding the low degree of differen-
tiation, which nowhere reaches the species level, the wild carp is a geo-
logically young species — dating at the most from the Pleistocene. From the
western and central Asian area of first occurrence and origin of the wild
carp, part of one strain later invaded the west, whereas part of another one
has penetrated to the east. Consequently, Cyprinus carpio carpio and C. c.
haematopterus exist today through their range as relatively poorly differen-
tiated taxa. The degree of relationship between the diverse strains of central
Asia and domestic stocks has yet to be established.
Dwarfed Wild Carp and Their Viability
Using wild adults selected on specially devised criteria (Misik and Tuca,
1965), morpho-ecological characters were studied first (Balon, 1958a,
1958b), and juveniles from the same spawn were used to evaluate the extent
of changes following culture in southern (Bastl, 1961 ) and northern (Rud-
zinski, 1961) European ponds (Slovakia-Poland). After developmental
studies were terminated, some of these juveniles were maintained in aquaria
in the laboratory and are discussed here. Though a more extensive study
was originally planned, as a result of personal circumstances only a part of
the data remained in my possession. In spite of this and the nine years
elapsed since the termination of this experiment a record of the results may
have some value.
Seven fish, hatched from eggs collected after carp spawning in a pond on
29 May 1956, were kept in small (30 1) aquaria. Each year, from 1956
through 1963, some of the experimental fish were preserved. From the others,
scales in the centre of body sides (key scales) were extracted, standard
lengths and weights were recorded, and the fish were returned to the aquaria.
The last fish was preserved after 7 years of continuous aquarium life. The
average temperature was 21 C C (range 18-24°C), although in the winter of
1958, as a result of a heating failure, the temperature was 10°C for several
Throughout the experimental period the fish were fed mainly with Tubijex
sp. at the rate of 1-2 g/week in the first 2 years and 3-4 g/week thereafter.
Only on a few occasions was the Tubijex replaced by the same weight of
scraps of beef or live Cladoccra. Although the diet was minimal to sustain
life, in the first 2 years it was sufficient for some growth. The explanation
could be that the ration provided, through that interval, some kind of nutrient
more suitable for the developmental steps in question (Balon, 1971a) than
for later steps, though even then the fish did not die as predicted by Vanccov
for such cases (Baton, 1960a ). During the 7 years of the experiment the fish
behaved normally, and gonads developed each year in the last four years,
although in the last five years the fish did not grow at all. These experiments
demonstrated the ability of the wild carp to survive on small rations of sub-
stitute food, the ability to survive starvation. (The term "starvation" is justi-
fied by the minimal maintenance diet given and by the remarkable changes
in body proportions of experimental fish.) As a consequence of starvation,
however, the ossification of some skeletal elements of starved carp sets in
earlier than in naturally developing carp (Balon, 1960b). The fish became
asymmetrically "dwarfed" due to artificially induced achondroplasia-likc
Starvation affected growth of experimental fish, which never exceeded the
average length and weight of the 1 -year-old fish from the natural habitat
(Table 1 ). Nonetheless, both wild and experimental animals became sexually
mature in the third growing season.
Growth of experimental fish was more than six-fold less than that of wild
fish in the Danube River. Furthermore, growth of experimental carp stopped
in the fourth year. Differences in growth are reflected in relative indices
(Table 1 ). The "index of average size" of the wild carp in the Danube was
7 and only 2 in experimental carp; the "index of weight growth" among wild
carp from a spawning school in the Danube River was 1052, whereas for
first generation reared in ponds it was 962, and among experimental fish,
Sexually-mature gonads of experimental carp were first found in a male
and female 34 months old preserved in March 1959. Ovaries of the female,
which measured 128 mm (standard length), weighed 0.5 g (representing
0.9% of the specimen's wet weight). The number of eggs was 3,950.
Ovaries of a female that was preserved 3 years later (in April 1962; age
5-f) weighed 2.8 g (left 1.56 g; right 1.16 g) and represented 5.8% of the
wet fish weight. The number of eggs, estimated by the same method (gravi-
metric), was 83,540. Ovaries of the last fish, a female that was 6-f years old
and which was preserved in January 1963, were thin, compact strips adher-
ing to the lateral and ventral sides of the intestine and contained large, single
eggs that were irregularly distributed. New gonadal tissue had begun to de-
velop around eggs from the previous year that had not yet been fully re-
sorbed. a phenomenon that is not unexpected for a fish unable to spawn
in an aquarium and forced to resorb the eggs for the last four years. Hence,
in these experimental fish, age at first attainment of sexual maturity and num-
ber of eggs produced are not related to size but to age. Did stress conditions
of starvation produce changes leading to a new type of stock dynamic
( Balon. 1 963b. p. 535 ) within the same generation?
Starving did bring forth distinct changes in body proportions (Fig. 2)
and in some meristic characters. Counts of fin rays of experimental fish are
within the range of those of wild carp (Misik, 1958) inhabiting the Danube
and those of the pond carp (Steffens, 1964). Scale counts and number of
gillrakers are also within the range of normal variability. Pharyngeal tooth
count and arrangement, however, are unusual (Table 2). Neither wild carp
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Fig. 2 Pharyngeal teeth of dwarfed wild carp no.
formula 1.3-3.1. approx. x8.
6. six-year-old female; tooth
TABLE 2. Morphometric and meristic characters of wild carp (Cyprinus
carpio carpio) from the Danube River and of its "dwarfed"
offspring from aquaria
Characters defined according to
' = 100
N = 7
% OF STANDARD LENGTH:
Length of head
Length of snout
Length of barbel i
Length of barbel n
Length of orbit
Postorbital length of head
Depth of head
Depth of body
Width of body
Length of caudal peduncle
Minimum depth of body
Pectoral origin to pelvic base
Pelvic base to anal base
Length of base of dorsal fin
Length of base of anal fin
Length of upper lobe of caudal fin
Length of lower lobe of caudal fin
Length of longest pectoral ray
Length of longest pelvic ray
Length of longest dorsal fin ray
Length of longest anal fin ray
% OF LENGTH OF HEAD:
Length of snout
Length of barbel i
Length of barbel n
Length of orbit
Postorbital length of head
Depth of head
Width of head
% OF LENGTH OF CAUDAL
Depth of caudal peduncle
Width of caudal peduncle
Minimum depth of body
",, OF PECTORAL ORIGIN TO
Length of longest pectoral ray
% OF PELVIC BASE TO ANAL BASE:
Length of longest pelvic ray
Characters defined according N = 100
N = 7
to MiSik, 1958
(ii) iii-iv 18-21 (22)
(n) hi 4-5
in (4) 5
iv-viii 16-18 iv-viii
vi-vii 17 (18) vi-viii
1(14) 15-18 (19)
I 8 (9)
Total number of scales
(34-36) 37-39 (40)
(35) 36, 38
in lateral series
(35-36) 37-39 (40)
(35) 36 (37)
Number of scales in
.' Number of gill-rakers
(22) 23-27 (28)
(22) 24, 26 (27)
on the first gill arch
(29) 30-34 (36)
1.2.1-3.1 11.3-3.1 1
* See figures on inside front cover.
nor domesticated pond carp had so many irregular teeth as experimental
carp (Fig. 2), though domestic pond carp show similar irregularities in 15%
more cases (Steffens, 1964) than wild carp. Dwarfed carp had considerably
longer heads, a longer first pair of barbels, larger eyes, longer preventral
distance, longer caudal fin lobes, and ventral fin rays (Table 2) than wild
carp from natural habitats. Dwarf carp were also characterized by a slightly
larger postorbital length of head, greater depth of head, greater predorsal
length, greater depth of body, greater length of the longest pectoral fin, and
longer ventral, dorsal, and anal fin rays in relation to the standard length. No
characters are considerably smaller in the dwarf fish except interorbital width
and postorbital length in relation to length of head. Length of snout, depth
and width of head, and length of the dorsal fin base are somewhat smaller
than those measurements in wild fish.
Similar differences in body proportions between wild and domestic carp
were found by Steffens ( 1964). Similar changes in morphometric characters
were described by Wunder (1949) for starved carp and for starved Euro-
pean bream (Abramis brama) by Liihmann and Mann (1957). In contrast,
Rudzinski (1961 ) reported that wild Danube carp in ponds had a smaller
head than a domestic carp of the type he studied.
The size of the gape of the mouth, a character that Rudzinski ( 1961 ) and
Steffens ( 1964) used so successfully to distinguish wild and domestic carps,
was considerably smaller in the dwarfed fish than in the domestic or wild
carp (Table 3). Rudzinski and Steffens considered the enlargement of the
mouth in domesticated animals to be an adaptation to changes in feeding
habits and also a probable result of artificial (man-induced) selectivity.
Domestic carp selectively adapted to utilize supplementary food added to
ponds grew better in ponds when artificial food was added. Is it possible
that the small amount of food given in small particles to dwarf carp of my
Fig. 3 Scale of dwarfed wild carp no. 6 with 5 annuli; the second annulus (for 1958)
is strongly intercepted, approx. xl6.
experiment may have brought about the development of a small gape of
mouth? The size of mouth would be an adaptive eharaeter in Kammerer's
(1923) sense then, in spite of criticism of such concepts (Kocstler, 1971).
The number of curves of the intestine in experimentally-dwarfed carp
varied (4,7,4,4,5,6, respectively) as did other characters (Table 3). Adult
domestic carp average six curves to the intestine (Klust, 1939). Hence not
all dwarf fish developed fewer curves. Rudzinski ( 1961 ) found, and StefTens
(1964) confirmed, that the intestine of wild carp was generally 15-25%
shorter than that of domestic carp. Artificially "dwarfed" carp developed the
shortest intestine which was less than 50% the length of that of the domestic
carp and slightly more than 50% of that of the wild carp. Differences are
even more emphasized if related to body weight: the ratio of gut length to
body weight is 2.2 in domestic carp, 3.0 in wild carp, and 8.3 in dwarfed
carp (Table 3).
In spite of experimental conditions of minimal temperature fluctuation,
the absence of seasonal changes, and constant feeding throughout the year.
Fig. 4 Scale of dwarfed carp no. 7, with absorbed dorsal and ventral edces, approx.
scales of dwarfed carp developed regular annual marks (Fig. 3). The acci-
dental drop in temperature in 1958 was indicated in the scales by a pro-
nounced annulus. The shape of most key scales was regular but heavy
absorption at the dorsal and ventral edges (Fig. 4) occurred in some col-
lected toward the front or rear end of the body during the last two years.
The first key scales were collected from the series above the lateral line,
dorsal to the origin of the pelvic fin. True annuli were readily distinguished,
but between them there were several confusing false annuli. I was, however,
unable to relate scale growth to body growth, and values of body size back-
calculated from scales for previous growth seasons do not make sense.
Interception of annuli may be initiated by the intervals of life history and
"the planetary yearly system" (Balon. 1971b. p. 99), but temperature or
other environmental factors may strengthen the annulus interception (Seger-
The above data are recorded as a stimulus to interest, and in order to
complete the description of the artificially dwarfed carp. The results of this
TABLE 3. Mouth-gape and intestine lengths indices for dwarfed, domestic
and wild carp: o/l = 10 X mouth-gape (cm-')/standard length
(cm), o/w = 10 X mouth-gape (cm-)/weight (g), o/lc = 10
X mouth-gape (cm' J )/length of head (cm); gut/1 = length of
intestine (tin) 1 cm of standard length, gut x w = length of
intestine (cm)/ 10 g of weight.
Dwarf wild carp
experiment extended conclusions of an earlier study (Balon, 1960b) beyond
the early intervals of fish life. Karzinkin's (1935) experiments suggested
that piscivorous fishes would be less capable of survival on substitute foods,
although some species, at least, produce dwarfed morphs (Popova, 1967)
if population survival is endangered. Some species that forage, however,
may not only survive when starved but if later given an optimum diet over-
take well-fed individuals (Krizenccky and Kfizenecka-Pulankova, 1953;
Kuznccov, 1957). Great viability is a characteristic of wild carp, which toler-
ate a wide range of environmental factors and arc consequently a unique
telcost species — a most successful colonizer of the world.
Probable Origin of Domestication
The most western natural occurrence of rhcophilic wild carp in the Danube
is at the mouth of the Morava (March) River. It was there, near Dcvin
(Toben), that the well-known Amber Road crossed the Danube River. The
Celtic settlements on the Danube River and the lower parts of its tributaries
in southeastern Slovakia "existed until the middle of the first century a.d. and
sometimes earlier (compare the settlement at Dcvin in the second half of
the first century), for the Latin material culture often mixed with the Dacian"
(translated from Pelikan, 1960). The carp was known to the Romans, who
travelled along the Amber Road through territory where huge schools of
wild carp probably spawned every year on Hooded meadows. Both facts indi-
cate that the Danube and Amber Road intersection is the site from which
transfer of the carp to European waters outside the Danube area probably
began. Other places are less probable, even though some were suggested.
Movtchan ( 1966) even assumed that Romans transported carp to Italy from
Fanagoria (the area of Kuban) and maintained them in ponds. Possibly,
again, Dubravius (1547) was confused by the transfer of carp to brackish
ponds and by its subsequent multiple spawning, which is, however, usual
for carp in warm water (Busehkiel, 1932). Conseqently, Dubravius' inter-
pretation of Pliny's (Plinius, a.d. 24-79; 1635) data concerning multiple
spawning of carp in Italy was incorrect and distracted his attention from
the true geographic origin of domestic carp.
ROMAN GOURMETS AND FIRST CARP IN CAPTIVITY
Although fish were captured in the Danube 10,000 years ago by the
Cromagnons of Upper Paleolite (France), the first proof of fishing activity
came from excavations of late Neolithic Stone Age (Clark, 1948). Childe
(1929) assumed that ". . . important are the deer's horn harpoons from
Vinca and Csoka (fig. 17), for they show how much the inhabitants of these
sites appreciated the good fishing of the Danube." When the Romans arrived
they encountered a well-developed fishing technique which was sufficiently
documented in the first writings on fishing in the Danube River, scattered
through the works of Strabo (63 B.C.; 1917-1932), Plinius (a.d. 24-79;
1635). Aelianus (a.d. 170-235; 1858), and Ausonius (a.d. 310-393;
1933). About a.d. 15 the Romans built a camp opposite the mouth of the
Morava River on the site of a Celtic village that later became their biggest
Pannonic town, Carnuntum, an important resting place on the Amber Road.
Stone reliefs from that period frequently used the fish motif, which I had in
two cases identified as a carp (e.g.. Fig. 5, Komarno City Museum). In the
first century a.d. the XlVth and XVth Roman legions were stationed on the
site of the present city of Bratislava, then called Peiso-Piso. Other garrisons
and troops, living at sites along nearly the entire length of the Danube River,
certainly fished for their livelihood as can be gathered from written remarks
made by Roman authors spoken of earlier, as well as from works of Herod-
Fig. 5 Stone relief with carp, partly spoiled by later sculptured head of a goat, from
the Roman excavations at Komarom (Komarno City Museum). Photo E. K.
otus (485-425 B.C.; 1920-1924), Paterculus Velleius (19 b.c.-a.d. 31;
1924), Tacitus (a.d. 55; 1836), and Cassiodorus (a.d. 490-585; 1626).
The commanders of these garrisons were members of patrician families and
as such were brought up on the Roman tradition of epicurianism. It must,
therefore, have been a source of pride to bring back to Rome an as yet un-
known delicacy. In Pliny's Historiae Naturalis, for example, are found such
Et in Danubio Mario extrahitur, And in the Danube River is fished
porculo marino similimus . . . , hausen (Huso huso) resembling a
sea dolphin . . . ,
and in Aelianus (loc. cit.) we find detailed descriptions of fishing through
ice. As early as the first century B.C. Cicero's teacher of gastronomy, Sergius
Orata, had devised special salt water reservoirs separated from the sea where
he stored fish for the kitchen. These reservoirs ensured a permanent supply
without regard to weather and success in fishing. According to Plinius (A.D.
24-79; 1635) this method was adopted by Lucinius Muraena, who began
storing freshwater fish. The patricians liked this manner of keeping their fish
and competed in establishing such "piscinae," often spending enormous sums
of money on them. Consul Lucullus (75 B.C.). whose reputation as a gourmet
is well known, dug through a hill near Naples to bring water to his ponds,
which were reputedly more costly than his villa.
Although the original idea of the piscinae, as devised by Sergius Orata,
was to store fish for gastronomic purposes, rearing fish later became a hobby.
Another patrician, Hortensius, became as famous as Lucullus, but for the
love of his ponds (Hortensius liked his eel so much that he wept when he
found it dead . . .), and was even accused by Cicero of neglecting politics
because of his fish. The Roman preferred sea fishes, as Varro (1 16-27 B.C.:
1912) emphasized; freshwater ponds were apparently considered inferior
and plebeian, but documented prejudice is at least proof of the existence of
Now perhaps it can be understood why visiting senators, patricians, and
plebeian soldiers not only tasted the fish during their temporary stay in the
Danube region but tried to transfer them alive to piscinae at home, either as
fish fanciers or as gourmet attractions for guests. Which of the largest, tastiest
fishes of this region would have survived the rigours of primitive transport?
Which could have lived in saline or brackish water after such transport? The
wild carp is among the least sensitive of fish and can tolerate water with a
low oxygen content so that it may be easily transported and afterwards be
kept for weeks in small reservoirs (transportation in wet moss or other
moisture-retaining materials is probable). Moreover, no special food is
needed and it can endure long periods of starvation. Besides having palatable
meat, carp has one very important ability — it lives in waters with salinities as
great as 6,000 mg Cl/I (Nakamura, 1948; Johnson, 1954; Mark, 1966).
Wild carp even live in the brackish waters of the delta of the Danube (Bana-
The above, then, is how I envisage the Danube wild carp to have been
transferred to Italy at the beginning of our era to begin life in piscinae. From
there some escaped to local rivers and. alter the collapse of the Empire and
the establishment of Christianity, were reared in monastery ponds. I do not
exclude the possibility that wild carp endemic to the Danube River were later
brought to other west European ponds. Apparently the messages of Roman
and Middle Age scholars describing these early imports were passed on from
generation to generation and probably accounted for the reestablishment of
the carp-rearing tradition. In support of the above let me quote Cassiodorus
. . . destinet carpam Danuvius: A ... from the Danube come carp
Rheno veniat anchorago (...): and from the Rhine herring. To pro-
sapori pisces de diversis finibus af- vide a variety of flavours, it is neces-
ferantur. sic decet regem pascere, ut sary to have many fish from many
a legatis gentium credatur paene countries. A king's reign should be
omnia possidere. such as to indicate that he possesses
Apparently King Theodorus (a.d. 490-526) of Ravenna (Italy) ordered the
transport of carp from the Danube to his country, thus imitating Roman
tastes and continuing the still novel introductions.
It was only 100 years ago that Dubisch brought Danube wild carp to the
Upper Vistula River for hybridization with the local form of domestic carp
(see Morcinek, 1909). In 1957 I repeated Dubisch's experiment by trans-
ferring 1.000 yearling Danube wild carp to the ponds at Ochaby (experi-
mental station of the Polish Academy of Sciences) (Rudzinsky, 1961 ). But
as in experiments on the Danube, studies at the Ochaby pond proved (Lesz-
czynska and Biniakowski. 1967) that wild carp are more suitable for stocking
into natural waters than are domestic carp.
FASTING MONKS REDISCOVER THE CARP
Christianity introduced more than 100 fasting days a year, and the only meats
that could be eaten during those days were crayfish, molluscs, and other cold-
blooded animals such as fish (in some regions fowl and/or unborn rabbit
embryos, laurices, were exceptions). As heavy punishment, which sometimes
went as far as the death penalty, was meted for violations of proscriptions, a
readily available supply of fish was important. Thus, fish became the only
possible foods for monks during a substantial part of the year. Although fish
were abundant in rivers and lakes, there were many days in the year when
weather conditions, natural disasters, or wars made fishing impossible.
Monks and missionaries, who had to follow fasting regulations, had difficulties
finding fish (Leonhardt, 1906). Probably after monks settled and established
a monastery, they thought about keeping a fish supply as they had in their
southern homeland; they also remembered or read how suitable the carp was.
The first monasteries were founded in the early 6th century (e.g., Monte
Cassino Monastery was founded a.d. 529 ) . Later they gained land and farms,
and conditions became favourable for the beginning of mass culture of fish
that provided the monastery a ready supply of food for fast days. Monks,
according to Leonhardt (1906). first reared local fishes: inland monasteries
Fig. 6 This illustration from the 18th century shows that monks, very much like the
Romans, enjoyed fishing for carp in monastery ponds. (Courtesy of the
Mansell Collection, London.)
kept pike (Esox lucius), crucian carp (Carassius carassius) , bream (Abramis
brama), and other species. But these species were difficult to keep in good
condition in the primitive reservoirs. They knew that carp could be bred
easily in these circumstances and introduced it.
REARING OF CARP IN PONDS AND FIRST REPRODUCTION IN CAPTIVITY
Carp then were first reared with other species of fish in a simple man-made
rearing pond. Certainly, some unexpected spawning occurred already in
Roman piscinae and also in monastery ponds; no organized reproduction,
however, was recorded. Charlemagne (a.d. 768-814), the first Holy Roman
Emperor, ordered his tenant farmers to maintain ponds and issued orders
for their control. His orders, however, were concerned with protection against
poaching, regulation of fishing and sale of fish, not with culture. Leonhardt
(1906) claims that the lack of concern in carp reproduction was conditioned
by an abundant fish fauna in local waters with which the landlords could
regularly stock the ponds. In my opinion, Leonhardt (loc. cit.) incorrectly
assumes the natural occurrence of the wild carp then in waters of southern
and northern Europe. Moreover, archaeological findings of Slavonic settle-
ments at the outset of the Polish Empire in the Ninth to Twelfth Centuries
did not include carp, though they have produced remains of ide (Leuciscus
idus), sturgeon (Acipenser sturio), chub (Leuciscus cephalus), tench (Tinea
tinea), perch (Perca fluviatilis) , mud loach [Misgurnus fossilis), roach
(Rutilus rutilus), beaked carp (Chondrostoma nasus). wels (Silurus glanis),
pike (Esox lucius), eel (Ani>uilla anguilla), salmon (Sahno salar or Sabno
trutta) and rapfen (Aspius aspius) (Perlbach, 1881 ). The first written evi-
dence of carp in Polish territory is dated 1466 and originates from the area
of Kotomyje in the Black Sea Basin (Chmielewski, 1965). Later carp were
kept in ponds as proved by the following words of M. Rej (a.d. 1505-1569)
quoted by Gorzynski ( 1964) :
... a wszystko to bardzo malem . . . and it is possible to obtain
zachodem otrzymac mozna: bo sie- everything without trouble: release
dem. ale dziewi^c karpi puscic, seven or nine carp, then the same
takiez w druga (sadzawke) karas- number into another pond of cru-
kow, ali ty i pieniazki i pozytek z cian carps, and you will benefit
tego mice mozesz. from it as well as have money.
But the pond culture of carp still cannot be connected with its pond repro-
Hildegarde ( 1089-1 170), mother superior of the Bavarian Convent of the
Benedictine Order, mentioned in her recipes the preparation of carp (Koch,
1925); Albertus Magnus (11937-1280; 1861) first wrote about breeding it
in ponds. But generally not much information is available about the rearing
of carp in the Middle Ages. It is as if the secret had been kept within the
monastery walls. Gradually, however, complete articles, even simple studies
about the rearing of the carp and its culture in ponds appeared (Dubravius,
1547; Strumieriski, 1573; Strojnowski, 1609; von Hohberg, 1687), the rein-
statement of rearing in the late Middle Ages is well known (Leonhardt, 1906;
Steffens, 1958). Von Hohberg (1687) considered that the rearing methods
of his time were superior to those used by the Romans and also that the
Romans would surely have enjoyed the taste of the real contemporary carp.
In the work of Dubravius (1547) numerous remarks about the beginning of
Roman domestication of carp piece together the historical connections.
ONE MORE ORIGIN OF DOMESTICATION
Domestication of wild carp in China began independently, occurred five
centuries earlier than in Europe (Leonhardt, 1906; Tamura, 1961; Hickling,
1962), and probably involved the local subspecies Cyprinus carpio haema-
topterus. The latest, anonymous Chinese study (the following quotations are
from the Russian translation — Anonymous, 1961) on pond culture stated
that "thanks to the creative efforts of the Chinese people for many genera-
tions, breeding of the carp in this country has proceeded successfully for
more than 2,000 .years. From China the breeding of this fish spread all over
the world." Further, it was Fan Lio (Tao Tschshugun) who succeeded in
spawning the carp and in growing the young to adulthood in a pond at the
time of the "Spring and Autumn" and the "Fighting Dynasties" (in the
eighth-third centuries B.C.). "From Asia the rearing of carp spread to
Europe and later to America, Australia, and Africa. " There is no reason for
the Chinese today to contradict an opinion that supports their national pride,
particularly as most European authors supported the Chinese origin of the
carp's domestication. The Chinese arc proud of their past isolation and inde-
pendent cultural development. As China was virtually inaccessible until a
century ago, would it not be improbable for the carp to have been brought
from there in the first years of the Gregorian calendar or the beginning of the
Middle Ages? An independent, second introduction of carp to Western
Europe probably occurred during the Middle Ages and is considered by some
to have been from China (Rudzinski, 1962). Previously mentioned evidence
favours the Danube as a source also of the later introductions. As a result of
the introduction into Asia of European pond carp, however, the carp in ponds
in some regions in eastern Asia may be mixed in origin (Buschkiel, 1933).
Consequences of Domestication
As early as the sixteenth century, races of carp were recognized, especially
those morphs in which scales were few or absent. The domestic carp of
western Europe was introduced into North America in 1831 for the first time,
again in 1872, and frequently later (Hessel, 1881; Bartlett, 1901, 1905;
Cole, 1905; Dymond, 1955; Atton, 1959; McCrimmon, 1968). The carp
was introduced into Australia in 1860 (Butcher, 1962, 1967), and in 1896
to the Cape of Africa (Jubb, 1967). According to my hypothesis all these
carp originated from the rheophilic wild carp of the Danube River. While
this hypothesis is geographically and historically attractive, in the absence of
more evidence it remains still speculative. With this in mind, the gold carp
(Cyprinus auratus) of the Schuykill and Massachusetts rivers (Forester,
1850, quoted by McCrimmon, 1968) could be a scaled carp whose genotype
was predominantly the wild morph and so selected a more lotic habitat than
other carp in the shipment whose genes were primarily of more advanced
domestic carp. (The most freshly caught Danubian wild carp that 1 saw were
always a clear golden colour on the scaled region of their bodies.)
In Europe the domestic carp became the most widely cultured and con-
sumed fish, and pond culture gradually became one of the sophisticated
branches of agriculture. The same species in North America, however, is
considered to be undesirable, probably because of its unpalatable meat as
compared to that of many local fishes and because of an adverse interaction
with the aquatic habitat. Even in Australia "The European carp (Cyprinus
carpio) has been proclaimed by legislation to be a 'noxious fish' and the
keeping and/or release of this species is prohibited" (Butcher, 1967).
Consequences of domestication of wild carp become apparent when the
wild morph from the Danube is compared with any established domestic
morph. Subsequent to rediscovery of rheophilic wild carp in the Czecho-
slovak-Hungarian section of the Danube (Balon and Misik, 1956) and
initial studies on it (Balon, 1957, 1958a; Misik, 1958), morphomctric and
growth indices were compared with those of the domestic populations (Tuca,
1958; Steffens, 1959; Bastl, 1961, 1962; Rudzinski, 1961; Chytra ct al.,
1961 ; and Misik and Tuca, 1965). Some morphological differences between
wild and domestic carp were stressed by Rudzinski (1961) and Steffens
( 1 964 ) . It will be sufficient to mention some of the results of the latter study :
The body of the domestic carp, which is nearly always much deeper than
that of the oblong, cylindrical body of the wild carp, appears to have more
flesh, but the calculated ratio of muscle in both morphs is the same. Without
regarding changes "in proportions, dressed weight of individual domesticated
carp did not increase even though its greater growth rate produces more
absolute meat in a given period of time. Probably the faster growth of domcs-
tic carp can be correlated with the larger mouth and longer intestines acquired
by the pond form following its adaptation to complementary nutrition and
perhaps also by man's selection.
The greater strength, mobility, and viability of wild carp are emphasized
by physiological characteristics. The wild carp has 189c to 19% more eryth-
rocytes and haemoglobin than docs the domestic carp. Blood sugar level is
16%— 26% higher. The wild carp has a much lower water content in muscles
and liver than does the domestic carp. Furthermore the wild carp has a
greater content of fat in individual organs, of glycogen in the liver, and of
vitamin A in the intestines, eyes, and liver. Consequently the taste of the wild
carp is better because the flesh is juicier. The wild carp is more mobile,
stronger, and nimbler because its muscles are better supplied with blood
ensuring a better supply of nutrients and elimination of waste; muscles do
not fatigue as quickly as do those of the domestic carp, enabling wild carp to
overcome the river current. The same improves the taste of the flesh of the
wild hare and the wild rabbit, said to be more juicy than the domestic rabbit
Epilogue The impressive number of these
that are 'not known in the wild state'
emphasizes the drastic nature of the
morphological changes that domes-
tication has so often brought about
in the transition from wild ancestor
to cultivate crop.
David R. Harris (1967, p. 91)
Perhaps the unpalatable taste of carp meat in North America is a result of
the carp's origin. Nearly all carp that live in natural waters here are descend-
ants of domestic carp which escaped into natural habitats. Limited ability to
utilize natural food and poor growth may have resulted in unpalatable flesh
in feral domestic carp. European domestic cattle, when left to become wild,
are known to remain in poor condition and never revert to a condition
similar to that of the strong, wild, ancestral auroch (Bos primigenius) (see
Talbot et al., 1965; Taylor. 1970. 1972). Perhaps the carp fall to feral con-
dition parallelled that of the cattle. If this is the case it is not surprising that
in North America carp "have never been wholeheartedly accepted by the
general public as an item of food" (Crossman, 1 969. p. 1 45 ) .
The need for fasting food may have been the main reason for rediscovery
of the carp in the Middle Ages, but not everywhere has the hedonistic
approach of the Romans to rearing of the carp given way to pure necessity;
there is some evidence (Fig. 6) of monks enjoying fishing for carp (Burton
and Burton. 1 968 ) . Nevertheless, the need for a larger variety of foods during
fasts undoubtedly played an important role in bringing about the tradition of
carp eating among some European Christians. Though related to fasting, this
tradition was eventually also embraced by some European Jews who made
the carp their Sabbath meal.
That the history of the domestication of the carp is generally similar to
that of the European rabbit (Oryctolagus cuniculus) may be of some interest
as peripheral support of my hypothesis on carp origin. Varro (1 16-27 B.C.;
1912) wrote that Romans brought rabbits from Spain and bred them in
special enclosures called leporaria to ensure a fresh supply of meat at all
times. Unborn rabbits collected from pregnant females were prepared as a
special dish called laurices. At the beginning of the Middle Ages rabbits were
a popular fare in western European monasteries, and laurices were eaten
even during fasting periods. (It was in the corridors and paved courts of the
monasteries that the rabbits began giving birth to their young above ground
instead of in burrows. The young gradually became domesticated and accus-
tomed to people.) In the middle of the sixteenth century several colour
morphs of rabbits were known. An albino rabbit is shown in Titian's
"Madonna" (1530) at the Louvre (Valcanover, 1960). Around 550 B.C.
the Chinese philosopher Confucius (1898) suggested to poor farmers that
they keep some rodents similar to rabbits (Volf, 1965). Unlike speculation
on the origin of the carp, the origin of the domestic rabbit was not assumed
to be in China. Otherwise the similarity of the domestic history of both
animals is striking.
Wild rabbits are overabundant in many countries of the world and conse-
quently special measures are taken to control them. Conversely, rheophilic
wild carp in Europe today is in danger of extinction. Regulation of the flow
of rivers and construction of waterworks arc destroying remaining spawning
areas. In the Danube River, between the mouths of the Morava and Hron
Rivers, spawning schools that were plentiful as recently as 15 years ago have
greatly decreased. At a decreased population density the wild carp may lose
its habitat to the domestic carp, which is repeatedly stocked, purposefully
and accidentally, into rivers. Competitive replacement may be accelerated if
planned hydroelectric plants and waterways are introduced (Balon, 1967a,
Scientifically directed selection would have a better chance of improving
cattle production and quality if genes of the ancestral auroch were still
available. The wild carp may be compared to the extinct auroch when
consequences of its possible extinction are contemplated. In 1965 Slovakia
included spawning schools of wild carp in the official list of animals protected
by law (Randik, 1967). Even so, because of the rapid deterioration of the
environment, extinction is the probable fate of this species. Protection of the
original stock of wild carp in the foothills of the Danube River is therefore
an important task for mankind. It was probably here that the most useful
domestic fish of the world originated.
If preglacial remnants of the carp from pile dwellings are rejected as incor-
rectly dated, it is postulated that the wild carp originated in central Asia and
spread naturally east into China and adjacent regions and west as far as the
Danube River in the last postglacial period. That this did not happen earlier
is suggested by the fact that in the Pleistocene, after the retreat of the conti-
nental glaciers, C. carpio was not among the species which penetrated either
Scandinavia, the British Isles, or North America. Had it occurred at that time
in eastern Asia it probably would have invaded North America together with
the pike and other freshwater fishes via the Bering land bridge.
The natural occurrence of the wild carp in the Danube River is inferred
from the different names given to the carp by the local inhabitants and from
the use of the same Celtic name outside the western limit of its natural range;
further evidence comes from medieval chronicles concerning the occurrence
of this fish, and from contemporary studies of the rheophilic wild form of the
carp in the Danube River. The ancient differentiation of wild carp in central
Asia and presence of undifferentiated isolated single morphs in the eastern
and western regions where it occurs suggest again that its place of origin must
be sought in central Asia.
A separate section is devoted to an experiment testing the ability of Danube
River wild carp to survive in starvation condition; this demonstrates the
hardiness of the animal and explains why it was domesticated for human
Finally, from written notes of the Romans and excavated art objects de-
picting the carp, the theory is presented that the first carp brought to Roman
piscinae came from places where the Amber Road — the most travelled thor-
oughfare of those times — crossed the Danube River. These localities, regu-
larly inundated meadows where schools of carp spawned, occurred at the
mouth of the Morava River, near ancient Carnuntum and the later Devin
Castle. Patrician commanders of the Roman legions stationed there brought
the carp to Rome as an epicurian delight, for it was one of the most hardy
species of fish and survived primitive techniques of transport and rearing.
After the decline and fall of the Roman Empire, the rearing techniques of
carp in Roman piscinae were not forgotten. Christian missionaries and monks
needed ready reserves of fish for their fasting days. From Latin writings and
from traditions of their southern homelands they knew that the carp was
appropriate for such purposes and looked after their supply as soon as they
settled somewhere. After the Middle Ages breeding had become routine, not
only in monasteries but also on private estates.
The history of the domestication of the carp in Europe that began with
wild carp from the Danube River (Cyprinus carpio carpio) is divided into
five periods: 1. First introduction by the Romans west of the Danube River
in the first to fourth century a.d.; 2. sporadic western introduction and rear-
ing in the fifth and sixth centuries; 3. beginning of rearing on a mass scale,
with some renewed introductions, and initial attempts to breed it in greater
quantities in specially-built ponds in the seventh to thirteenth centuries;
4. beginning of breeding, mass culture, and fortuitous selection in the four-
teenth to sixteenth centuries; 5. intensification of breeding, purposeful selec-
tion, and introductions into North America. Australia, the Far East, and
Africa from the seventeenth century. In China domestication of the carp,
probably from local subspecies C.c. haematopterus, was begun five cen-
turies earlier but independently of similar efforts in Europe.
The most important features by which the wild carp from the Danube
River differ from the present domesticated forms are stressed and reference
is made to a striking similarity with the domestication of the rabbit. Attention
is drawn to the necessity of preserving the last wild schools of this fish in the
foothill-zone of the Danube River and of protecting the wild predecessor of
the domestic carp and its habitat. Most important, a repetition of the fate of
the extinct auroch, the ancestor of cattle, should not be permitted. As with
cattle, domestic carp in wild conditions becomes an animal of poor food
quality, which is perhaps the reason why the carp deteriorated so after its
Above all, I owe an outstanding debt of gratitude to V. Misik, V. Tuca, I.
Bastl (Slovak Agricultural Academy, Bratislava) and E. Rudziriski (Polish
Academy of Sciences, Cracow) for their original studies of the Danube wild
carp that were consistent with my ideas and which led to the formulation of
the theorem here presented. I thank J. Holcik (Slovak National Museum,
Bratislava), O. Oliva (Charles University. Prague) and A. G. Coche
(UNDP/FAO, Chilanga), who pointed out errors or inadequacies in the
first draft, and E. J. Crossman (Royal Ontario Museum, Toronto) for read-
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VOOREN, C. M.
1972 Ecological aspects of the introduction of fish species into natural habitats
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no. 4, pp. 565-583.
1949 Fortschrittliche Karpfenteichwirtschaft. Stuttgart.
1925 Tritt der Karpfen schon im Diluvium Norddeutschlands auf ? Mitteilg. d.
Fischereivereine Brandenburg-Pommern etc., vol. 17, pp. 80-83.
P e 3 K) m e
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meMy cnoco6HOCTb ahkoto Kapna peKH JXynaPi Bbi-
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Aenne Kapna 6i»ijio y>Ke pyTHHoft ne TOJibKO b MOHa-
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HcTopnH pa3Be^.enHH Kapna b EBpone, na i iiiiiaio-
uiancn jxuKHM KapnoM H3 peKH .Hynan (Cyprinus carpio
carpio), ac/ihtch na nHTb nepHO^OB: 1 nepno/i, b
kotopom nepBbiMH ikhuuih pa3Bo^nTb Kapna Phm-
;hihc, Ha 3ana,n. ot peKH JXyuavi, b nepBOM n flo neT-
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necKoro iianajia pa3Be,ueHHfl Kapna b iihtom h inec-
tom cTOJieTHJix; 3 - nepnoA MaccoBoro pa3Be,aeHini
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niecTHaAHaToe ctojicthh; 5 nepnoA ycujieHiioro
pa3BeAeiiHH, ucieycTpeMJieHHan ce.ieKunn 11 nanajio
pa3BeAeniiH Kapna b CeBepnoii AMcpiiKe, ABCTpa.iHH,
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jieTHH. Bo3mo>kho b KuTac pa3Be,aeHne Kapna naMa-
;h)ci» ii3 MecTHoro noABH^a C. c. haematopterus 500
.ieT paHbiue, ho He 38Bhchmo ot Ebpoiii.i.
3;j.ocb noa i iepKiiiuieTCH ca&roe Baxuioe cbomctbo,
no KOTopoMv aiiKHii Kapn peKH /Iynai'i OTJiHiaeTCfl
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ccbunca Ha pajiiTo.'ihiioc cxoactbo MexcAy pa3Be,aenn-
om Kapna n npiipymiBaHiieM KpoJiHKa. Taione o6pa-
maeTcn BuiiMannc na noTpe6nocTb coxpanenHn noc-
.lejmix ahkhx craii Kapna b npe^ropbHx peKH Jlyuati
H npejoxpanenne jiiKoro npe^Ka KWibTiiBiipoBaHHoro
wapna b ero ecTecTBennou cpeae. II caMoe rjiaBHoe,
noBTopeHHe cyji.b6bi Bbiwepujero Typa, npe^Ka Kpyn-
hoto poraToro CKOTa, He aojdkho 6biTb AonycTHMo.
KaK KpynnbiH poraTbift ckot, TaK ii Ky.ibTHBHpoBaii-
HblH Kapn B AHKOM COCTOHHHH HB.IHeTCH pbl60H n.10-
xoro KawecTBa. 3to, bo3.\io>kho, h HB.iHeTCH npHMHHOH
-roro, mo Kapn cra;i He nony.inpubiM b AiuepHKe noc-
."ie ero hhtpciykuhh.